Adaptive forest management improves stand-level resilience of temperate forests under multiple stressors.

Forests are expected to be strongly affected by modifications in climate and disturbance regimes, threatening their ability to sustain the provision of essential services. Promoting drought-tolerant species or functionally diverse stands have recently emerged as management options to cope with global change. Our study aimed at evaluating the impact of contrasting stand-level management scenarios on the resilience of temperate forests in eastern North America and central-western Europe using the individual process-based model HETEROFOR. We simulated the evolution of eight stands over 100 years under a future extreme climate according to four management scenarios (business as usual - BAU; climate change adaptation - CC; functional diversity approach - FD; no management - NM) while facing multiple disturbances, resulting in a total of 160 simulations. We found that FD demonstrated the greatest resilience regarding transpiration and tree biomass, followed by CC and then BAU, while these three scenarios were equivalent concerning the net primary production. These results were however dependent on forest type: increasing functional diversity was a powerful option to increase the resilience of coniferous plantations whereas no clear differences between BAU and adaptive management scenarios were detected in broadleaved and mixed stands. The FD promoted a higher level of tree species diversity than any other scenario, and all scenarios of management were similar regarding the amount of harvested wood. The NM always showed the lowest resilience, demonstrating that forest management could be an important tool to mitigate adverse effects of global change. Our study highlighted that tree-level process-based models are a relevant tool to identify suitable management options for adapting forests to global change provided that model limitations are considered, and that alternative management options, particularly those based on functional diversity, are promising and should be promoted from now on.

A harmonized database of European forest simulations under climate change.

Process-based forest models combine biological, physical, and chemical process understanding to simulate forest dynamics as an emergent property of the system. As such, they are valuable tools to investigate the effects of climate change on forest ecosystems. Specifically, they allow testing of hypotheses regarding long-term ecosystem dynamics and provide means to assess the impacts of climate scenarios on future forest development. As a consequence, numerous local-scale simulation studies have been conducted over the past decades to assess the impacts of climate change on forests. These studies apply the best available models tailored to local conditions, parameterized and evaluated by local experts. However, this treasure trove of knowledge on climate change responses remains underexplored to date, as a consistent and harmonized dataset of local model simulations is missing. Here, our objectives were (i) to compile existing local simulations on forest development under climate change in Europe in a common database, (ii) to harmonize them to a common suite of output variables, and (iii) to provide a standardized vector of auxiliary environmental variables for each simulated location to aid subsequent investigations. Our dataset of European stand- and landscape-level forest simulations contains over 1.1 million simulation runs representing 135 million simulation years for more than 13,000 unique locations spread across Europe. The data were harmonized to consistently describe forest development in terms of stand structure (dominant height), composition (dominant species, admixed species), and functioning (leaf area index). Auxiliary variables provided include consistent daily climate information (temperature, precipitation, radiation, vapor pressure deficit) as well as information on local site conditions (soil depth, soil physical properties, soil water holding capacity, plant-available nitrogen). The present dataset facilitates analyses across models and locations, with the aim to better harness the valuable information contained in local simulations for large-scale policy support, and for fostering a deeper understanding of the effects of climate change on forest ecosystems in Europe.

Evidence, causes, and consequences of declining nitrogen availability in terrestrial ecosystems.

The productivity of ecosystems and their capacity to support life depends on access to reactive nitrogen (N). Over the past century, humans have more than doubled the global supply of reactive N through industrial and agricultural activities. However, long-term records demonstrate that N availability is declining in many regions of the world. Reactive N inputs are not evenly distributed, and global changes-including elevated atmospheric carbon dioxide (CO2) levels and rising temperatures-are affecting ecosystem N supply relative to demand. Declining N availability is constraining primary productivity, contributing to lower leaf N concentrations, and reducing the quality of herbivore diets in many ecosystems. We outline the current state of knowledge about declining N availability and propose actions aimed at characterizing and responding to this emerging challenge.

Defoliation Change of European Beech (Fagus sylvatica L.) Depends on Previous Year Drought.

European beech (Fagus sylvatica L.) forests provide multiple essential ecosystem goods and services. The projected climatic conditions for the current century will significantly affect the vitality of European beech. The expected impact of climate change on forest ecosystems will be potentially stronger in southeast Europe than on the rest of the continent. Therefore, our aim was to use the long-term monitoring data of crown vitality indicators in Croatia to identify long-term trends, and to investigate the influence of current and previous year climate conditions and available site factors using defoliation (DEF) and defoliation change (ΔDEF) as response variables. The results reveal an increasing trend of DEF during the study period from 1996 to 2017. In contrast, no significant trend in annual ΔDEF was observed. The applied linear mixed effects models indicate a very strong influence of previous year drought on ΔDEF, while climate conditions have a weak or insignificant effect on DEF. The results suggest that site factors explain 25 to 30% DEF variance, while similar values of conditional and marginal R2 show a uniform influence of drought on ΔDEF. These results suggest that DEF represents the accumulated impact of location-specific stressful environmental conditions on tree vitality, while ΔDEF reflects intense stress and represents the current or recent status of tree vitality that could be more appropriate for analysing the effect of climate conditions on forest trees.

Simulating tree growth response to climate change in structurally diverse oak and beech forests.

This study aimed to simulate oak and beech forest growth under various scenarios of climate change and to evaluate how the forest response depends on site properties and particularly on stand characteristics using the individual process-based model HETEROFOR. First, this model was evaluated on a wide range of site conditions. We used data from 36 long-term forest monitoring plots to initialize, calibrate, and evaluate HETEROFOR. This evaluation showed that HETEROFOR predicts individual tree radial growth and height increment reasonably well under different growing conditions when evaluated on independent sites. In our simulations under constant CO2 concentration ([CO2]cst) for the 2071-2100 period, climate change induced a moderate net primary production (NPP) gain in continental and mountainous zones and no change in the oceanic zone. The NPP changes were negatively affected by air temperature during the vegetation period and by the annual rainfall decrease. To a lower extent, they were influenced by soil extractable water reserve and stand characteristics. These NPP changes were positively affected by longer vegetation periods and negatively by drought for beech and larger autotrophic respiration costs for oak. For both species, the NPP gain was much larger with rising CO2 concentration ([CO2]var) mainly due to the CO2 fertilisation effect. Even if the species composition and structure had a limited influence on the forest response to climate change, they explained a large part of the NPP variability (44% and 34% for [CO2]cst and [CO2]var, respectively) compared to the climate change scenario (5% and 29%) and the inter-annual climate variability (20% and 16%). This gives the forester the possibility to act on the productivity of broadleaved forests and prepare them for possible adverse effects of climate change by reinforcing their resilience.

The Effect of Environmental Factors on the Nutrition of European Beech (Fagus sylvatica L.) Varies with Defoliation.

Despite being adapted to a wide range of environmental conditions, the vitality of European beech is expected to be significantly affected by the projected effects of climate change, which we attempted to assess with foliar nutrition and crown defoliation, as two different, yet interlinked vitality indicators. Based on 28 beech plots of the ICP Forests Level I network, we set out to investigate the nutritional status of beech in Croatia, the relation of its defoliation and nutrient status, and the effects of environmental factors on this relation. The results indicate a generally satisfactory nutrition of common beech in Croatia. Links between defoliation and nutrition of beech are not very direct or very prominent; differences were observed only in some years and on limited number of plots. However, the applied multinomial logistic regression models show that environmental factors affect the relationship between defoliation and nutrition, as climate and altitude influence the occurrence of differences in foliar nutrition between defoliation categories.

Isotopic evidence for oligotrophication of terrestrial ecosystems.

Human societies depend on an Earth system that operates within a constrained range of nutrient availability, yet the recent trajectory of terrestrial nitrogen (N) availability is uncertain. Examining patterns of foliar N concentrations and isotope ratios (δ15N) from more than 43,000 samples acquired over 37 years, here we show that foliar N concentration declined by 9% and foliar δ15N declined by 0.6-1.6‰. Examining patterns across different climate spaces, foliar δ15N declined across the entire range of mean annual temperature and mean annual precipitation tested. These results suggest declines in N supply relative to plant demand at the global scale. In all, there are now multiple lines of evidence of declining N availability in many unfertilized terrestrial ecosystems, including declines in δ15N of tree rings and leaves from herbarium samples over the past 75-150 years. These patterns are consistent with the proposed consequences of elevated atmospheric carbon dioxide and longer growing seasons. These declines will limit future terrestrial carbon uptake and increase nutritional stress for herbivores.

The response of soil solution chemistry in European forests to decreasing acid deposition.

Acid deposition arising from sulphur (S) and nitrogen (N) emissions from fossil fuel combustion and agriculture has contributed to the acidification of terrestrial ecosystems in many regions globally. However, in Europe and North America, S deposition has greatly decreased in recent decades due to emissions controls. In this study, we assessed the response of soil solution chemistry in mineral horizons of European forests to these changes. Trends in pH, acid neutralizing capacity (ANC), major ions, total aluminium (Altot ) and dissolved organic carbon were determined for the period 1995-2012. Plots with at least 10 years of observations from the ICP Forests monitoring network were used. Trends were assessed for the upper mineral soil (10-20 cm, 104 plots) and subsoil (40-80 cm, 162 plots). There was a large decrease in the concentration of sulphate (SO42-) in soil solution; over a 10-year period (2000-2010), SO42- decreased by 52% at 10-20 cm and 40% at 40-80 cm. Nitrate was unchanged at 10-20 cm but decreased at 40-80 cm. The decrease in acid anions was accompanied by a large and significant decrease in the concentration of the nutrient base cations: calcium, magnesium and potassium (Bc = Ca2+  + Mg2+  + K+ ) and Altot over the entire dataset. The response of soil solution acidity was nonuniform. At 10-20 cm, ANC increased in acid-sensitive soils (base saturation ≤10%) indicating a recovery, but ANC decreased in soils with base saturation >10%. At 40-80 cm, ANC remained unchanged in acid-sensitive soils (base saturation ≤20%, pHCaCl2 ≤ 4.5) and decreased in better-buffered soils (base saturation >20%, pHCaCl2 > 4.5). In addition, the molar ratio of Bc to Altot either did not change or decreased. The results suggest a long-time lag between emission abatement and changes in soil solution acidity and underline the importance of long-term monitoring in evaluating ecosystem response to decreases in deposition.

Soil parent material-A major driver of plant nutrient limitations in terrestrial ecosystems.

Because the capability of terrestrial ecosystems to fix carbon is constrained by nutrient availability, understanding how nutrients limit plant growth is a key contemporary question. However, what drives nutrient limitations at global scale remains to be clarified. Using global data on plant growth, plant nutritive status, and soil fertility, we investigated to which extent soil parent materials explain nutrient limitations. We found that N limitation was not linked to soil parent materials, but was best explained by climate: ecosystems under harsh (i.e., cold and or dry) climates were more N-limited than ecosystems under more favourable climates. Contrary to N limitation, P limitation was not driven by climate, but by soil parent materials. The influence of soil parent materials was the result of the tight link between actual P pools of soils and physical-chemical properties (acidity, P richness) of soil parent materials. Some other ground-related factors (i.e., soil weathering stage, landform) had a noticeable influence on P limitation, but their role appeared to be relatively smaller than that of geology. The relative importance of N limitation versus P limitation was explained by a combination of climate and soil parent material: at global scale, N limitation became prominent with increasing climatic constraints, but this global trend was modulated at lower scales by the effect of parent materials on P limitation, particularly under climates favourable to biological activity. As compared with soil parent materials, atmospheric deposition had only a weak influence on the global distribution of actual nutrient limitation. Our work advances our understanding of the distribution of nutrient limitation at global scale. In particular, it stresses the need to take soil parent materials into account when investigating plant growth response to environment changes.

Forest soils in France are sequestering substantial amounts of carbon.

The aim of this study was to assess whether French forest soils are sources or sinks of carbon and to quantify changes in soil organic carbon (SOC) stocks over time by resampling soil in long-term forest monitoring plots. Within each plot, and for each survey, soils were sampled at five points selected in five subplots and divided into layers. Composite samples were produced for each layer and subplot, then analysed for mass, bulk density and SOC. Linear mixed models were used to estimate SOC changes over 15years between two soil surveys carried out in 102 plots in France. A factor analysis and a budget approach were also used to identify which factors and processes were primarily responsible for SOC dynamics. Forest soils throughout France substantially accumulated SOC (+0.35MgCha-1yr-1) between 1993 and 2012. The SOC sequestration rate declined with stand age and was affected by stand structure. Uneven-aged stands sequestered more SOC than did even-aged stands (p<0.001). For the forest floor, the SOC sequestration rate estimated by the budget approach was in agreement with that based on stock comparison. This increasing SOC stock in the forest floor can be explained by recent changes in certain factors affecting litter decomposition (climate and litter quality). For the mineral soil, the budget approach was unable to replicate the observed SOC sequestration rate, probably because SOC stocks were not yet at equilibrium with litter inputs at the beginning of the monitoring period (contrary to our steady-state assumption). This explanation is also supported by the fact that the SOC sequestration rate decreased with stand age. As the SOC sequestration rate declines with stand age and is higher in uneven-aged stands, forest management has the potential to influence this carbon sink.

Tree mineral nutrition is deteriorating in Europe.

The response of forest ecosystems to increased atmospheric CO2 is constrained by nutrient availability. It is thus crucial to account for nutrient limitation when studying the forest response to climate change. The objectives of this study were to describe the nutritional status of the main European tree species, to identify growth-limiting nutrients and to assess changes in tree nutrition during the past two decades. We analysed the foliar nutrition data collected during 1992-2009 on the intensive forest monitoring plots of the ICP Forests programme. Of the 22 significant temporal trends that were observed in foliar nutrient concentrations, 20 were decreasing and two were increasing. Some of these trends were alarming, among which the foliar P concentration in F. sylvatica, Q. Petraea and P. sylvestris that significantly deteriorated during 1992-2009. In Q. Petraea and P. sylvestris, the decrease in foliar P concentration was more pronounced on plots with low foliar P status, meaning that trees with latent P deficiency could become deficient in the near future. Increased tree productivity, possibly resulting from high N deposition and from the global increase in atmospheric CO2, has led to higher nutrient demand by trees. As the soil nutrient supply was not always sufficient to meet the demands of faster growing trees, this could partly explain the deterioration of tree mineral nutrition. The results suggest that when evaluating forest carbon storage capacity and when planning to reduce CO2 emissions by increasing use of wood biomass for bioenergy, it is crucial that nutrient limitations for forest growth are considered.

[Applicability analysis of spatially explicit model of leaf litter in evergreen broad-leaved forests].

The spatially explicit model of leaf litter can help to understand its dispersal process, which is very important to predict the distribution pattern of leaves on the surface of the earth. In this paper, the spatially explicit model of leaf litter was developed for 20 tree species using litter trap data from the mapped forest plot in an evergreen broad-leaved forest in Tiantong, Zhejiang Pro- vince, eastern China. Applicability of the model was analyzed. The model assumed an allometric equation between diameter at breast height (DBH) and leaf litter amount, and the leaf litter declined exponentially with the distance. Model parameters were estimated by the maximum likelihood method. Results showed that the predicted and measured leaf litter amounts were significantly correlated, but the prediction accuracies varied widely for the different tree species, averaging at 49.3% and ranging from 16.0% and 74.0%. Model qualities of tree species significantly correlated with the standard deviations of the leaf litter amount per trap, DBH of the tree species and the average leaf dry mass of tree species. There were several ways to improve the forecast precision of the model, such as installing the litterfall traps according to the distribution of the tree to cover the different classes of the DBH and distance apart from the parent trees, determining the optimal dispersal function of each tree species, and optimizing the existing dispersal function.

Biomass and nutrient content of sessile oak (Quercus petraea (Matt.) Liebl.) and beech (Fagus sylvatica L.) stem and branches in a mixed stand in southern Belgium.

Accurate estimates of the amounts of nutrients immobilised in the organs and tissues of different tree species are of prime importance to make appropriate tree species selection and determine the harvesting regime that will ensure forest sustainability. Sixteen sessile oaks (Quercus petraea (Matt.) Liebl.) (64-129years; stem diameters: 17-57cm) and twelve beeches (Fagus sylvatica L.) (43-86years; stem diameters: 9-50cm) were destructively sampled from a mixed stand located on an acid brown soil in southern Belgium. Statistical models were developed to investigate the differences in nutrient concentrations between tree species, between aboveground tree compartments of the same species, and between tissues of the same compartment. For stem tissues, vertical concentration profiles were described using a versatile equation. Allometric equations were used to predict biomass and nutrient content of tree compartments based on tree dimensions. Broadly speaking, nutrient concentrations tended to be somewhat higher for oak compared with beech, but the amplitude and the direction of inter-species differences varied greatly, depending on the nutrient and the tree compartment. For both species, living branch nutrient concentrations tended to decrease with increasing branch diameter, except for Ca (oak) and Mg (beech). Nutrient concentrations were consistently higher in bark than in wood; this difference between tissues was quite pronounced for Ca, particularly in the case of oak. The biomass and nutrient content equations were used to investigate the effects of tree species and harvesting regime on nutrient exports at harvesting. For equivalent harvesting scenarios, beech was found to induce higher Mg exports than oak, and inversely for Ca. Assuming stand clear cutting, complete tree harvesting would increase average nutrient exports from 65% (Ca) to 162% (P) compared with a stem-only harvesting scenario. These results provide valuable information in the current context of the more intensive utilization of forest products.

Spatial and temporal patterns of throughfall chemistry within a temperate mixed oak-beech stand.

The effects of meteorological variables and local canopy characteristics on throughfall chemistry were investigated within a mixed oak-beech stand during the leafed and the leafless seasons. Throughfall was collected after each rain event in 12 structural units delimited by three neighbouring trees each. For each investigated ion (H+, Mg2+, Ca2+, K+, NH4+, Na+, Cl(-), NO3(-) and SO4(2-)), mixed linear models were used to analyse the effects of the rain volume (R) and of the length of the preceding dry period (ADP) on net throughfall ion fluxes and to evaluate the influence of canopy density and tree species composition on these relationships. The models explained more than 70% of the total variability. Canopy characteristics accounted for most of the spatial variability while a large part of the temporal variability was not explained by R and ADP, suggesting an influence of other variables such as wind force and direction. For each season, net throughfall ion fluxes generally increased with increasing R and ADP, whose coefficients were interpreted as leaching capacity and dry deposition rate, respectively; H+ is an exception with negative exchange capacity (i.e. absorption) during the leafed season. Dry deposition rate and exchange capacity estimates were generally lower for the leafless canopies compared with the leafed season. For a given density level, exchange capacities were either higher for oak than for beech or similar for both species while dry deposition tended to increase with increasing beech proportion. The enhancing effect of canopy density on dry deposition was particularly marked for the mixed leafed canopies. For the pure canopies, the effect of canopy density on dry deposition rate and exchange capacity tended to be more pronounced during the leafless period compared with the leafed season, while it was generally limited compared with the species effect.

Effects of biological and meteorological factors on stemflow chemistry within a temperate mixed oak-beech stand.

The effects of meteorological variables and tree species on stemflow chemistry were investigated within a mixed oak-beech stand during the leafed and the leafless seasons. Stemflow was collected after each rain event. For each investigated ion (H(+), Mg(2+), Ca(2+), K(+), NH(4)(+), Na(+), Cl(-), NO(3)(-), SO(4)(2-)), mixed linear models were used to analyse the effects of the rain volume (R) and of the length of the preceding dry period (ADP) on net stemflow ion fluxes as well as to assess the effect of tree size on these relationships. The models generally explained more than 70% of the total variability. The product between trunk circumference and tree height (C130Htot) explained most of the inter-individual variability, except for oak during the leafed season for which the effect of tree size was not significant or very limited. On the other hand, besides R and ADP, other rain event characteristics like wind force and direction were suggested to also partly explain the inter-event variability. For each season, net stemflow ion fluxes tended to increase with increasing R and ADP, whose coefficients were interpreted as leaching and dry deposition rates, respectively; exceptions were negative exchange rates (i.e. absorption) for NH(4)(+) and NO(3)(-) during the leafless period and for H(+) during the leafed season. Moreover, when it appeared significant, the effect of tree size corresponded to an increase of leaching, absorption and/or dry deposition rates as C130Htot increased. Exchange rate estimates were generally higher for the leafless season compared with the leafed period as well as for beech compared with for oak. Dry deposition rate estimates were generally higher for the leafless season compared with the leafed period. Differences in dry deposition rates between both species were particularly pronounced for the leafless season with much higher estimates for beech compared with oak.

Modeling leaf dispersal in mixed hardwood forests using a ballistic approach.

In mixed-species stands, modeling leaf litter dispersal is important to predict the physical and chemical characteristics of the forest floor, which plays a major role in nutrient cycling and in plant population dynamics. In this study, a spatially explicit model of leaf litterfall was developed and compared with two other models. These three models were calibrated for a mixed forest of oak and beech using litterfall data from mapped forest plots. All models assumed that an allometric equation described individual leaf litter production, but they strongly differed in the modeling of the probability density of leaf shedding with distance from source trees. Two models used a negative exponential function to account for leaf dispersal with distance, and this function was allowed to vary according to wind direction in one of them. In contrast, our approach was based on a simple ballistic equation considering release height, wind speed, wind direction, and leaf fall velocity; the distributions of wind speeds and wind directions were modeled according to a Weibull and a Von Mises distribution, respectively. Using an independent validation data set, all three models provided predictions well correlated to measurements (r > 0.83); however, the two models with a direction-dependent component were slightly more accurate. In addition, parameter estimates of the ballistic model were in close agreement with a foliar litter production equation derived from the literature for beech and with wind characteristics measured during leaf litterfall for both species. Because of its mechanistic background, such a spatially explicit model might be incorporated as a litterfall module in larger models (nutrient cycling, plant population dynamics) or used to determine the manner in which patch size in mixed-species stands influences litter mixture.